tIS: IS and relatives with passenger genes
Over the past few years, a number of TE have
been identified which are very closely related to known IS but which carry
passenger genes not directly involved in transposition. These elements, simpler
than known Tns, are called transporter ISs (tISs) (Siguier, et al., 2009) (Fig
genes include transcription regulators (e.g. ISNha5, members of the IS1595 family), methyltransferases (e.g. IS220, IS1380 family), and antibiotic
resistance (e.g. ISCgl1, IS481family) genes and can be located
upstream, downstream or on both sides of the transposase gene (Fig 1.14.1). They can include a significant
amount of DNA with no clear coding capacity (e.g.ISBse1, ISSpo3 and ISSpo8, IS1595 family) and are longer than typical IS (e.g. ISCausp2,
7915 bp, IS1595 family). This has presumably delayed their
identification. Since the second IS end would occur at an unexpectedly distant position, they would resemble partial IS copies lacking a second end.
tISs are generally present in low copy number.
Many occur only in single copy in a given genome raising the question of
whether they are active. Moreover, more than one closely related but
non-identical derivative can be found in a single genome (e.g. ISSpo3, andISSpo8 in Silicibacter
pomeroyi DSS-3 and ISNwi4 and ISNha5 in Nitrobacter
winograski Nb-255). Others
are present in more than one copy. ISPre3,
an IS66 relative from Pseudomonas resinovorans plasmid pCAR1
includes a hypothetical protein and is present as 2 copies with different
insertion sites as judged by their typical 8 bp DR (Gourbeyre unpublished).
The mechanisms involved in acquisition of
additional genes to generate tISs are at present unclear. They do not appear to
carry programmed recombination systems as do some members of the Tn3 family. One possibility is that tIS
are derived by deletion from ancestral compound transposons. These are composed of two ISs flanking any DNA segment
either in direct or inverted orientation (see references in (Craig, et al.,
2002)). The flanking IS are able to mobilize the intervening DNA segment
(Fig 1.2.3). They were among of the first
types of transposon described and include the models Tn5 (flanking IS50) and Tn10 (flanking IS10) as well as Tn9 (flanking IS1). Since these early
examples, many other such composite transposons have been identified either by
experiment or from genome sequencing.
Tn number registry
Some early observations concerning Tn5 and Tn10 suggest that the flanking IS can undergo mutation rendering them
less autonomous (for example mutations within one IS which inactivate its
transposase; see (Mahillon & Chandler, 1998)).
Furthermore, studies on IS102 from
the pSC101 plasmid clearly indicated that transposition can occur using one
established IS end and a second surrogate end located at some distance from the
et al., 1982). If, as in this case, the intervening DNA includes a
passenger gene, this creates a novel transposon. It has also been observed that
other ISs such as IS911 can use
surrogate ends during transposition (Polard, et al., 1994). Moreover,
isolated individual IS ends are often observed in sequenced genomes and could
provide a source of surrogate ends. A number of potential transposable elements
of this type, containing a single IS, a passenger gene and a short flanking
sequence resembling an additional IS end, have been observed in nature. These
include: colistin resistance (MCR-1) (Liu, et al.,
2016) which is
associated with ISApl1 (Susu He pers.
Comm.), the fabI gene of Staphylococcus haemolyticus resulting in triclosan resistance and
associated with IS1272 and the blaCTX-M-19 gene associated with ISEcp1B (Poirel, et al.,
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Wang Y, Walsh TR, et al. (2016)
Emergence of plasmid-mediated colistin resistance mechanism MCR-1 in animals
and human beings in China: a microbiological and molecular biological study. Lancet Infect Dis 16: 161-168.
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L, Decousser JW & Nordmann P (2003) Insertion sequence ISEcp1B is involved
in expression and mobilization of a bla(CTX-M) beta-lactamase gene. Antimicrob Agents Chemother 47: 2938-2945.
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M (2009) The new IS1595 family, its relation to IS1 and the frontier between
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